Journal of the Royal Society of Western , 96: 41–53, 2013

Avifauna of the Mulga–Eucalypt Line in

D T BELL 1,*, R C BELL 1 & W A LONERAGAN 2

1 5 The Rise, Mindarie, WA 6030, Australia. 2 School of Plant Biology, University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia. * Corresponding author  [email protected]

The Mulga–Eucalypt Line has long been recognised as a vegetational transition from Acacia spp. domination in the northeast to Eucalyptus spp. domination in the southwest. The avifauna of this transition zone was sampled over four seasons at four locations representing a north to south geographic gradient of ~170 km. Each location comprised one survey site dominated by acacias and a second dominated by eucalypts. A total of 81 avian were recorded over the July 2011 to April 2012 sampling period with an average sample richness of 19 species per plot and a mean plot density of 1.05 /ha. Most species were classed as sedentary (39%), with the remainder as locally dispersive (21%), nomadic (25%) or migratory (15%). Ordination analysis indicated a latitudinal gradient of species from north to south along the first axis associated with gradually increasing rainfall. The second ordination axis revealed avian species differences dependent on the dominant vegetation. The third ordination axis displayed differences in bird species related to season, especially the spring and summer samples. The avifauna assemblage structure of northern locations was strongly influenced by the inclusion of irruptive, arid-zone nomadic species. Site richness comparisons between locations were not different, but eucalypt- dominated sites had more species than acacia-dominated sites. Site richness was greater during spring and summer. Density also was influenced by the influx of arid-zone nomadic species to the northern sites of the transition zone during spring and summer. Species diversity of the samples was influenced by vegetation type, but the measures of evenness and dominance were not affected by the factors of location, vegetation type or season. The Mulga–Eucalypt Line transition zone presently has considerable conservation value, but is under threat from pressures of agricultural and mining intrusions. Continued protection of native vegetation in the conservation reserves should be encouraged.

KEYWORDS: avifauna community, density, gradient analysis, Mulga–Eucalypt Line, seasonal variation, species richness, vegetation.

INTRODUCTION mesic Mediterranean climate of the southwest with wet winters and dry summers. Within the transition zone, The Mulga–Eucalypt Line is a zone of vegetation change mulga woodlands with trees 3–4 m in height occur on running inland from the Western Australian coastal town shallow, red-loam soils overlying hardpan (Beard 1990; of Geraldton southeastward through Menzies in the Patrick 2001). Eucalypt woodlands in this region with Goldfields (Beard 1990). The line separates the Eremaean trees to 10 m occur on deeper loam soils with the depth Botanical Province to the north and the South-West to the hardpan favouring different eucalypt species. Botanical Province to the south (Figure 1) and also separates the Southwestern and the Central Provinces of Birds of arid-zone mulga regions of Western Australia the Western Faunal Sub-Region (Kikkawa & Pearse have been surveyed by Cody (1994), Recher & Davis 1969). (1997), Burbidge et al. (2000, 2010) and Bell et al. (2013). Serventy & Whittell (1962) noted that bird populations The predominant tree species of the arid regions to the along the Mulga–Eucalypt Line were an admixture of northeast is mulga (), associated with a birds of the more arid woodlands, scrublands and number of other acacias (e.g. A. pruinocarpa, A. linophylla, hummock grasslands to the north and east and those of A. coriacea and A. acuminata). The predominant tree the more mesic woodlands and forests to the south and species of the more mesic areas to the southwest are west. A number of studies of the avifauna of Eucalyptus- eucalypts (e.g. Eucalyptus salmonophloia, E. loxophleba, E. dominated sites south of the Mulga–Eucalypt Line have salubris and E. transcontinentalis) (Boland et al. 1984). The been published (Ford 1971; Davies 1977; Arnold & vegetation transition from acacia to eucalypt domination Weeldenburg 1998; Recher & Davis 2002, 2010; Bell et al. has been variously attributed to climatic variables (Beard 2007, 2010). Bell et al. (2007) showed that there was a 1990), predominately rainfall (Patrick 2001), although gradient of avian assemblage change from northeast to edaphic parameters play a role within the transition zone southwest from surveys of exclusively eucalypt (Beard 1990). The Mulga–Eucalypt Line roughly lies woodlands in the northern part of the South-West between the 250–350 mm rainfall isohyets and separates Botanical Province. A study specifically designed to the arid zone of the northeast with limited rainfall determine if the avifauna changes across the Mulga– occurring in both summer and winter, from the more Eucalypt Line or if birds inhabiting Acacia-dominated habitats are different from those of adjacent Eucalyptus- dominated habitats in the transition zone has not been © Royal Society of Western Australia 2013 attempted. The objectives of this study were to: (i)

41 Journal of the Royal Society of Western Australia, 96(2), December 2013

METHODS

Study sites The avifauna of the Mulga–Eucalypt Line was sampled at four locations (Barnong Nature Reserve, Lochada Nature Reserve, Charles Darwin Reserve and Buntine Nature Reserve) (Figure 1). The four sampling locations also represented a north to south geographic gradient of ~170 km. Details of the geographic location of each study site and the dominant plant species are presented in Table 1. All sites are in native vegetation regions managed for conservation and have been unburned for many years. The Lochada sites are in the Karara and Lochada Important Bird Area (IBA) and the Charles Darwin sites are in the Mt Gibson and Charles Darwin IBA (Dutson et al. 2009). Mean annual rainfalls for the nearest Bureau of Meteorological Stations (www.bom.gov.au/climate) for each site were 261 mm for Barnong (Yalgoo), 291 mm for Lochada (Morawa), 282 mm for Charles Darwin (Paynes Find) and 342 mm for Buntine (Buntine).

Sampling procedures Each of the four sampling locations included two avian survey sites, one in Acacia-dominated vegetation and a second in Eucalyptus-dominated vegetation. The survey site vegetation, however, was not completely exclusive, since the understory of the Eucalyptus-dominated sites had a number of shrubby acacias and the Acacia- dominated sampling locations had sporadic eucalypt trees. Each permanently marked survey site was a plot of 500 m radius (78.54 ha) centred in a region of Figure 1 Study area map depicting northwestern region representative vegetation and divided into four of the Mulga–Eucalypt Line vegetation transition zone quadrants (0-90o, 90-180o, 180-270o and 270-360o). The first between the Eremaean Botanical Province in the two authors shared the field observation and recording northeast and the South-West Botanical Province in the of data. Each quadrant was traversed during a half-hour southwest with the local towns in black circles and the period and all birds heard or sighted were identified, study sites in black diamonds. tallied and recorded. Two periods of sampling, one in the early morning and one in the late afternoon, were made for each quadrant in each vegetation type for each document the avifauna of this vegetation zone; (ii) season. All four quadrants for a single site were surveyed determine if a gradient in the avifauna occurs from north consecutively. Sampling took place in mid-winter (3–14 to south across this region; (iii) assess whether there are July 2011), mid-spring (10–22 September 2011), mid- differences in bird assemblages from sites dominated by summer (30 December 2011 to 7 January 2012) and mid- acacia versus nearby sites dominated by eucalypts; and autumn (10–18 April 2012). No samples were taken (iv) determine possible seasonal differences in bird during rainy or excessively windy conditions. There were assemblages. no major eucalypt-flowering episodes at any of the

Table 1 Mulga–Eucalypt Line sampling site information.

Location Latitude Longitude Major species

Barnong acacia 28°25.65'S 116°07.86'E Acacia linophylla, Melaleuca uncinata Barnong eucalypt 28°25.40'S 116°08.52'E Eucalyptus loxophleba, E. oleosa Lochada acacia 29°12.04'S 116°30.89'E A. aneura, A. linophylla Lochada eucalypt 29°12.14'S 116°31.53'E E. salmonophloia, E. loxophleba Darwin acacia 29°35.14'S 116°57.57'E A. aneura, Callitris spp., Melaleuca spp. Darwin eucalypt 29°36.62'S 116°55.13'E E. salmonopholia, E. loxophleba Buntine acacia 29°58.02'S 116°35.29'E A. acuminata, Casuarina spp. Buntine eucalypt 29°58.80'S 116°34.74'E E. loxophleba

42 Bell et al.: Birds of the Mulga–Eucalypt Line sampling sites or seasons. All sites in a single location measures ANOVA was used to test aspects of the avian were sampled at one time, but the order of the locations assemblage structure, richness, density, species diversity, was randomised in each season’s surveys. Sampling was evenness and dominance (dependent variables) against also randomised for vegetation type and then the independent variables geographic location, vegetation particular sampling quadrant. type and seasonal variation (Zar 1996). Where the Avian nomenclature followed Christidis & Boles ANOVA indicated significance Holm-Sidak post-hoc (2008). Movement status designations were determined multiple comparison calculations were made to compare from the works of Saunders & Ingham (1995), Johnstone differences between groups. No significant interactions & Storr (1998, 2004), Morcombe (2000), Simpson & Day were found between the main factors in any ANOVA. (2004), Recher & Davis (2002, 2010) and Bell et al. (2007, While the original data were found to be not normally 2010). distributed, even after square-root transformation, all data met the equal-variance test. The results reported are based on square-root transformed data. All statistical Statistical analyses testing was carried out using Sigma Stat® 3.5 (Systat Relative density of each species in each sample was Software Inc. 2006). calculated by dividing the number of observations of that species by the total number of birds observed in the sample. In order to reveal patterns of overall avian RESULTS community structure and patterns of individual bird species distributions, the matrix of relative density data General avian community was analysed by Detrended Correspondence Analysis A total of 81 species were recorded in the 2011–2012 (DCA) (Jongman et al. 1995). The matrix included only samples of birds of the Mulga–Eucalypt Line region, bird species that occurred in at least two samples. including 30 non- and 51 passerines Characteristics of the avian population of each site (Appendices 1, 2). Species richness values for the were also described in terms of site richness, density and individual sample locations were similar, ranging species population heterogenity. Avifauna richness was between site totals of 47 and 54 species. Species totals determined as the total number of bird species recorded separated by vegetation type were also similar with 68 at each site during the 4 hours of sampling for each species recorded in Acacia-dominated sites and 66 species season. Site avifauna density (birds/ha) was the total in Eucalyptus-dominated sites (Table 2). Of the species number of observations of all birds for the two surveys recorded, 39% were classed as sedentary, 21% as locally (am and pm) divided by two and then by 78.54 (sample dispersive, 25% as nomadic and 15% as migratory. Red- area site). Due to the major irruption of the population of capped robin and chestnut-rumped thornbill were the budgerigars in the spring, counts continued only up to a most frequently observed species, each occurring in 31 of maximum of 100 individuals within any one quadrant. the 32 separate samples. Other sedentary and/or locally- As a result, bird densities during spring could have been dispersive species occurring in more than 75% of samples underestimated. Three measures of site species included the Australian raven, Australian ringneck, grey heterogeneity were determined: Shannon indices of shrike-thrush and inland thornbill. diversity (H’ = Σp log p), evenness (J’ = H’/H’ ) and i 10 i max Nomadic and/or migratory species appeared in the the Simpson index of dominance (D = Σp 2), where p i i records of particular seasons. Among these highly mobile equals the proportion for the ith species. These indices species, which were observed only in winter and spring, combine both species richness and the evenness of were the black-eared, pallid and Horsfield’s bronze distribution of individuals within the sample (Peet 1974). cuckoos, regent parrot and white-winged triller. Black The Shannon indices are biased toward species richness, and pied honeyeaters and the rufous were only while the Simpson index of dominance is biased toward recorded in the spring samples. Species restricted to only the density of the most common species (Magurran 1988). the spring and summer samples included budgerigar, Relationships between geographic location, vegetation cockatiel, red-backed kingfisher, black-faced cuckoo- type and seasonal variation and the DCA site axis scores shrike and white-fronted honeyeater. Rainbow bee-eater, were explored by correlation analysis and where and crimson chat were recorded only in significant associations found, differences were tested by the summer and zebra finch was recorded only in unpaired t-tests (Zar 1996). A two-way repeated summer and autumn.

Table 2 Species richness and movement type percentages of the avifauna of the Mulga–Eucalypt Line including all samples, samples of acacia sites alone and eucalypt sites alone and the separate seasons alone.

All Acacia Eucalypt Winter Spring Summer Autumn

Richness 81 68 66 49 57 53 38 Movement Type Sedentary 39% 38% 39% 49% 37% 43% 47% Locally Dispersive 21% 22% 21% 20% 18% 27% 26% Nomadic 26% 25% 24% 17% 28% 21% 16% Migratory 14% 15% 15% 14% 17% 9% 11%

43 Journal of the Royal Society of Western Australia, 96(2), December 2013

Avian assemblage patterns Particularly diagnostic species (those with very high The DCA ordination procedure utilised a matrix of 62 or very low DCA species scores) influenced the position avian species and 32 samples, after singletons were of the samples along the geographic gradient of the first removed from the complete set of relative densities. The DCA axis (Appendices 1, 2). Northern species included first DCA axis explained 48% of the sample variance and black and pied honeyeaters, budgerigar, cockatiel, zebra separated the samples on a geographic gradient (Figure finch and little button-quail. All these arid-zone birds 2). The highest first axis sample scores were from were classed as nomadic or migratory birds. Species with Barnong. Lochada samples tended to be grouped in the low DCA axis 1 values were in the more southerly and central section of the first axis and Darwin and Buntine mesic end of the geographic gradient and included the samples were located on the low-score end of the first black-faced cuckoo-shrike, black-faced woodswallow, grey fantail, yellow-rumped thornbill and red wattlebird. axis. Correlation analysis revealed a significant (r30 = 0.61, p ≤0.05) relationship between sample latitude and the Avian species strongly influencing the second DCA DCA first axis score. sample axis and related to vegetation differences The second DCA axis, which explained another 22% included the singing honeyeater, black-faced cuckoo- of the sample variance, separated samples from the shrike, rufous songlark, crested pigeon and mulga parrot acacia sites at the higher score end of DCA axis 2 from which had higher scores and were associated with acacia the eucalypt site scores at the lower end (Figures 2, 3). sites. Species with lower scores that were associated An unpaired t-test of the second axis DCA sample scores primarily with samples from eucalypt sites included the between the acacia sites and the eucalypt sites indicated rufous treecreeper, red-backed kingfisher, weebill, regent parrot, striated pardalote and Major Mitchell’s cockatoo. that the bird assemblages were significantly different (t30 = 4.59, p ≤0.001) between vegetation types. Budgerigar, cockatiel, western corella and crimson The third DCA axis explained a further 15% of the chat, species recorded in greatest numbers in spring, sample variance and separated the spring samples at the influenced the samples on upper end of DCA sample upper end of axis 3 from the summer samples at the axis 3. Species with low DCA species axis 3 scores were lower end of axis 3 (Figure 3). One-way analysis of rainbow bee-eater, Nankeen kestrel and Australian variance for axis 3 sample scores indicated that season magpie, which were recorded only in summer. ≤ also had a significant (F3,28 = 3.95, p 0.05) effect on the bird assemblages of the region.

Figure 2 Detrended Correspondence Analysis sample plot for axis 1 values against axis 2 values. The first two letters of the name of the site, the vegetation type and the sampling season designate each sample. On axis 1, values related to the samples from the same location are enclosed by lines of different types: Barnong –––––––; Lochada –– –– ––; Charles Darwin – – –; Buntine ····. On axis 2, values related to the two vegetation types are separated by the dash-dot-dash line.

44 Bell et al.: Birds of the Mulga–Eucalypt Line

Avian community characteristics sample richness between the four locations, the mean Overall, sample richness averaged 19.5±0.8 species per number of bird species in eucalypt sites was significantly ≤ sample (Table 3). However, there was considerable greater than in acacia sites (F1,3 = 30.75, p 0.01). The variation between samples. The lowest number of species repeated measures ANOVA showed richness of samples in a single sample was 14 recorded during winter at the in spring and summer to be significantly greater than the ≤ Barnong acacia site. The highest number of species samples of winter and autumn (F3,9 = 6.90, p 0.01). recorded was 28 at the Barnong eucalypt site during Bird density in the Mulga–Eucalypt Line during 2011– summer. Although there were no statistical differences in 2012 was generally low at a mean of 1.05 birds/ha, but

Table 3 Richness (no. of species), density (birds/ha), species diversity (H’), evenness (J’) and dominance (D) of samples of the Mulga–Eucalypt Line. Values with same superscript within the same factor are not different at p ≤0.05.

Richness Density Diversity Evenness Dominance

Overall 19.47 ± 0.76 1.05 ± 0.19 1.03 ± 0.02 0.80 ± 0.02 0.15 ± 0.02 Location Barnong 18.88 ± 2.00 a 1.79 ± 0.69 a 0.89 ± 0.10 a 0.71 ± 0.08 a 0.23 ± 0.09 a Lochada 20.50 ± 1.57 a 0.97 ± 0.08 a 1.08 ± 0.04 a 0.83 ± 0.02 a 0.11 ± 0.01 a Darwin 19.50 ± 1.50 a 0.87 ± 0.19 a 1.04 ± 0.03 a 0.81 ± 0.02 a 0.11 ± 0.02 a Buntine 19.00 ± 1.18 a 0.58 ± 0.06 a 1.10 ± 0.07 a 0.87 ± 0.04 a 0.16 ± 0.04 a Vegetation Acacia 17.56 ± 1.09 a 0.82 ± 0.20 a 0.99 ± 0.04 a 0.80 ± 0.03 a 0.15 ± 0.03 a Eucalyptus 21.38 ± 1.07 b 1.29 ± 0.30 b 1.07 ± 0.06 b 0.81 ± 0.04 a 0.16 ± 0.04 a Season Winter 16.88 ± 1.03 ac 0.61 ± 0.09 a 1.00 ± 0.03 a 0.82 ± 0.01 a 0.15 ± 0.01 a Spring 22.50 ± 1.29 b 1.91 ± 0.70 a 1.00 ± 0.13 a 0.74 ± 0.08 a 0.28 ± 0.09 a Summer 21.88 ± 1.42 bc 0.93 ± 0.09 a 1.11 ± 0.04 a 0.83 ± 0.02 a 0.11 ± 0.02 a Autumn 16.63 ± 1.31 a 0.77 ± 0.17 a 1.00 ± 0.04 a 0.82 ± 0.02 a 0.07 ± 0.01 a

Figure 3 Detrended Correspondence Analysis sample plot for axis 2 values against axis 3 values. The first two letters of the name of the site, the vegetation type and the sampling season designate each sample. On axis 2, values related to the two vegetation types are separated by dash-dot-dash line as in Figure 2. On axis 3, values related to the samples from the same seasonal samples are enclosed by lines of different types: Autumn –––––––; Spring –– –– ––; Winter– – –; Summer ····.

45 Journal of the Royal Society of Western Australia, 96(2), December 2013 highly variable (Table 3). Only vegetation type had a associated with Acacia-dominated sites of the region. All ≤ significant (F1,3 = 16.06, p 0.05) influence on the mean these species have been previously associated density of birds in the study samples, with eucalypt sites predominantly with Acacia spp. (although not strictly having greater density compared to acacia sites. Acacia aneura-dominated vegetation sites) (Cody 1994; The Shannon measure of species diversity (H’) Burbidge 2010; Bell et al. 2013). Species more likely to be averaged 1.03±0.02 overall (Table 3). As with the associated with Eucalyptus-dominated woodlands measures of bird density, only samples varying by included the rufous treecreeper, weebill, red-backed ≤ kingfisher, tawny frogmouth, Major Mitchell’s cockatoo, vegetation type were significantly (F1,3 = 10.34, p 0.05) different with eucalypt-dominated sites being more red-tailed black cockatoo, regent parrot, brown goshawk diverse than acacia-dominated sites. The statistical and brown falcon. The differential use of particular tree comparisons of the evenness (J’) and dominance (D) species by birds recorded in our study has been observed showed no difference due to location, vegetation type or in a range of other studies (Hartley 1953; Franzreb 1978; season of samples. Although there were no statistical Noske 1979; Ford et al. 1986; Holmes & Robinson 1981). differences in sample richness between the four locations, A number of reasons have been suggested for the the repeated measures ANOVA (square-root transferred) preference of particular tree species by birds. The showed the mean number of bird species in eucalypt sites abundance, biomass and availability of resources and the relative abundance of particular tree species in a was significantly greater than in acacia sites (F1,3 = 30.75, p ≤0.01); furthermore, the richness of samples in spring region may have relevance to insectivorous birds and summer to be significantly greater than the samples choosing certain tree species (Abbott & Van Heurck of winter and autumn (F = 6.90, p ≤0.01). 1985). Acacia and Eucalyptus support entirely different 3,9 psyllid subfamilies (New 1988), but data from studies from woodlands and forests in Western Australia indicate that there is high overlap among the DISCUSSION eaten by bird species (Calver & Wooller 1981; Wooller & Calver 1981; Tullis et al. 1982; Abbott & Van Heurck 1985; During the sampling period of July 2011 to April 2012, Carver et al. 1991; Majer et al. 1997). Leaf morphological the avifauna of the Mulga–Eucalypt Line varied in and nutrient differences in leaves between tree species relation to a north to south geographic gradient mainly may also be relevant in facilitating the search for insects in the pool of available sedentary and locally dispersive among foliage elements (Jackson 1979; Braithwaite et al. species, but also as a result of the presence of a number 1989; Recher et al. 1996; Watson 2011). Large trees for of nomadic and migratory birds in northern sites. There nesting and perching may be related to the presence of was a secondary influence of vegetation type, with some particular raptors (Storr 1984; Aumann 2001). birds associated with Acacia-dominated sites and others hollows for some species of parrots could also influence a associated with Eucalyptus-dominated sites. Seasonal preference for eucalypt-dominated sites. variation in the avifauna of sites appeared small, as the majority of species were either sedentary or locally Seasonal differences dispersive species. Minor seasonal differences were due Minor differences in avifauna assemblage of the Mulga– to the influx of nomadic and migratory species during Eucalypt Line were also found between the four seasonal spring and summer. samples of the July 2011 to March 2012 sampling period. As 70% of the bird species recorded were classed either Geographic pattern as sedentary of locally dispersive, seasonal differences The geographic variation in avian assemblage noted in might be expected to be small. However, a wide number the Mulga–Eucalypt Line has been recorded for other of arid zone birds in Australia are known to be highly Western Australian regions. A biogeographic gradient in responsive to changes in available food and water avian population structure associated primarily with resources (Davies 1984; Pavey & Nano 2009). Bell et al. rainfall was documented along the northern portion of (2007) found that the influx of nomadic honeyeaters the South-West Botanical Province in vegetation during episodes of eucalypt flowering strongly dominated by eucalypt species (Bell et al. 2007). Also, a influenced the avian associations of eucalypt sites to the gradual, broad-scale change in avian community south of the Mulga–Eucalypt Line. In the current study, structure due to climatic factors was the primary black and pied honeyeaters, two opportunistic influence in a study of biogeographic patterns in birds of nectarivores, occurred in the northern sampling sites the Carnarvon Basin, well to the north of the present during spring, but a general lack of flowering episodes study region (Burbidge et al. 2000). Birds in the southern in eucalypt species during the sampling period could sites of the study area were primarily sedentary and have limited any variation in avian assemblage structure locally dispersive species, while the more northern sites due to season. Three species of migratory cuckoos, included a number of arid-zone nomadic species. Horsfield’s bronze cuckoo, pallid cuckoo and black-eared cuckoo, were recorded in the Mulga–Eucalypt Line samples, but none were numerous. Other Mulga– Influence of vegetation type Eucalypt Line species listed as migratory, including the The samples of the bird assemblages of the Mulga– striated pardalote, white-winged triller, rufous songlark, Eucalypt Line also showed differences in composition tree martin and grey fantail, were also recorded between the Acacia-dominated sites and nearby occasionally during the 2011–2012 sampling period, but Eucalyptus-dominated sites. The singing honeyeater, were never numerous. Nix (1976) characterised a number black-faced cuckoo-shrike, rufous songlark, crested of Western Australian species as migratory. Keast (1968) pigeon and mulga parrot were species strongly also reported information related to the seasonal

46 Bell et al.: Birds of the Mulga–Eucalypt Line migratory movements of a range species, including a Bird density of sites of the Mulga–Eucalypt Line number of honeyeaters. It is apparent, however, that averaged 1.05±1.07 birds/ha. However, during spring the further analysis of the movement patterns of arid-zone sample in the Eucalyptus-dominated site of the Barnong birds and the classification of the species as either Nature Reserve rose to 5.87 birds/ha due primarily to the migratory or nomadic is needed. The lack of tagging and irruption in populations of budgerigars and cockatiels. recapture studies of terrestrial birds of Australia is a The density of birds in sites of the Mulga–Eucalypt Line major impediment in the determination of the potential was generally much lower than in more mesic Australian influence of migration on the presence of species in arid- habitats. Density of birds in tropical eucalypt savanna zone samples (Griffioen & Clarke 2002). woodlands of the Northern Territory is quite high at 9.4 birds/ha (Woinarski & Tidemann 1991). Density of birds Bird richness, density and heterogeneity in southwestern Western Australian jarrah forest is also relatively high at 5.2 birds/ha (Nichols & Muir 1989). In Mean species richness within the Mulga–Eucalypt Line mallee-boombush (Melaleuca uncinata) habitats of was not affected by location, although richness of northeastern Victoria, bird density is ~3.0 birds/ha eucalyptus sites was greater than acacia sites and sample (Gilmore 1985). Bird density in relation to habitat richness in spring and summer was enhanced over resource availability obviously requires further research. samples from autumn and winter. Generally, habitats that are more productive, of greater maturity and/or have Species diversity (H’) in the Mulga–Eucalypt Line greater habitat heterogenity are richer in averaged 1.03. Evenness (J’) averaged 0.80 and dominance assemblages (Ricklefs & Schluter 1993). However, this (D) averaged 0.15. Species diversity values of the winter does not seem the case for Western Australian bird sample Eucalyptus-dominated sites of Bell et al. (2007) in assemblages, as samples from a very wide range of sites the more mesic region to the south of the Mulga–Eucalypt all have similar site richness values. The mean richness Line averaged slightly higher at 1.12, possibly due to the of samples of the avifauna of the Mulga–Eucalypt Line at more even distribution of the densities of individuals 19.5 species was similar to those recorded for avifauna of among the species. The measures of heterogeneity in the the Pilbara much further to the north, as measured by Mulga–Eucalypt Line, however, were quite variable, Burbidge et al. (2010), which averaged 19.1 species. In the probably due to the influx of a number of irruptive species Doolgunna and Mooloogool Rangelands of the of the arid interior during spring and summer. Our northeastern Gascoyne, also to the north of the present findings generally concur with those of Keast (1985), who samples, species richness averaged 21.7 species per stated that eucalypt communities throughout Australia sample (Bell et al. 2013). Managed eucalypt forest sites, vary little in total avian species diversity, although the well to the southwest of the Mulga–Eucalypt Line, causes are not known. typically contain 10–33 bird species per site (Williams et All sites utilised in this study were within protected al. 2001; Abbott et al. 2003). Eucalyptus-dominated sites in conservation reserves, although much of the Mulga– the northern regions of the South-West Botanical Eucalypt Line region is increasingly being affected by Province, also south of the present study, averaged 27.3 mining, wheat farming and sheep-grazing activities. species per sample in the study by Bell et al. (2007). Continued protection of the reserves by the Western Cousin & Phillips (2008) hypothesised that scarcity of Australia Department of Parks and Wildlife and food resources results in a species richness threshold independent conservation organisations, such as Bush beyond which there is insufficient resources to support Heritage Australia, is essential for the maintenance and additional species as habitat complexity increases. Low conservation of the avifauna of this important vegetation species richness of Western Australian regions relative to transition zone between the arid zone of central Australia other Australian regions has also been associated with and the woodlands and forests of the southwest Western reduced numbers of nectar-feeding species and fewer Australia. litter-, trunk- and bark-gleaning species (Nichols & Muir 1989; Woinarski et al. 1997). The lack of eucalypt flowering episodes during our sampling period might ACKNOWLEDGEMENTS have further influenced possible richness differences due We appreciate the chance to sample the avifauna on the to season by reducing the number of expected nectar- Charles Darwin Reserve, owned and administered by feeding nomadic honeyeaters. Careful documentation of Bush Heritage Australia. Dale Fuller, Resident Manager vegetation structural complexity could provide more of the Charles Darwin Reserve, was especially helpful. conclusive evidence to compare species richness values The other sampling locations are under the management between north to south gradients in Western Australia of the Western Australia Department of Park and and to compare the species richness of Western Australia Wildlife (DPaW). Statistical advice was kindly provided sites to comparably complex habitats of eastern Australia. by Matt Williams (DPaW) and Pieter Poot and Michael Slight changes in the avifauna of spring and summer Renton, both of the School of Plant Biology, University of in the samples of the Mulga–Eucalypt Line were mainly Western Australia. Ian Abbott and Michael Craig due to the inclusion of a few nomadic and migratory provided valuable insights to improve the presentation species to the sedentary and locally dispersive species of the final manuscript. present in all seasons. Reduced impetus for migration due to the general lack of a large seasonal variation in resource availability has previously been highlighted for REFERENCES the restricted richness of Western Australian avifaunal ABBOTT I, MELLICAN A, CRAIG M D, WILLIAMS M, LIDDELOW G & assemblages (Nichols & Muir 1989) and may be an WHEELER I 2003. Short-term logging and burning impacts on influence affecting the richness of samples in our study. species richness, abundance and community structure of

47 Journal of the Royal Society of Western Australia, 96(2), December 2013

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Dell B, Havel J J & Malajczuk N (eds) The jarrah forest, pp. DAVIES S J J F 1977. Man’s activities and bird’s distribution in the 133–153. Kluwer Academic Publishers, Dordrecht. arid zone. Emu 77, 169-172. NIX H A 1976. Environmental control of breeding, post-breeding DAVIES S J J F 1984. Nomadism as a response to desert conditions dispersal and migration of birds in the Australian region. In: in Australia. Journal of Arid Environments 7, 183–195. Frith H J & Calaby J H (eds) Proceedings of the XVI DUTSON G, GARNETT S & GOLE C 2009. Australia’s important bird International Ornithological Congress, pp. 272–305. Australian areas—key sites for bird conservation. Birds Australia RAOU Academy of Science, Canberra. Conservation Statement 15. Birds Australia, Carlton. NOSKE R A 1979. Coexistence of three species of treecreepers in FORD J 1971. Distribution and taxonomy of southern birds in the north-eastern New South Wales. Emu 79, 120–128. Great Victorian Desert. Emu 71, 27–36. 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PAVEY C R & NANO C E M 2009. Bird assemblages of arid SIMPSON K & DAY N 2004. Field guide to the birds of Australia (7th Australia: vegetation patterns have a greater effect than edition). Penguin Books Australia Ltd., Ringwood. disturbance and resource pulses. Journal of Arid Environments STORR G M 1984. Birds of the Pilbara Region, Western Australia. 73, 634–642. Records of the Western Australian Museum, Supplement 16, 1– PEET R K 1974. The measurement of species diversity. Annual 63. Review of Ecology and Systematics 5, 285–307. TULLIS K J, CALVER M C & WOOLLER R D 1982. The invertebrate RECHER H F & DAVIS W E 1997. The foraging ecology of a mulga diets of small birds in Banksia woodland near Perth, W.A. bird community. Wildlife Research 24, 27–43. during winter. Australian Wildlife Research 9, 303–309. RECHER H F & DAVIS W E 2002. Foraging profile of a Salmon WATSON D M 2011. A productivity-based explanation for Gum woodland avifauna in Western Australia. Journal of the woodland bird declines: poorer soils yield less food. Emu Royal Society of Western Australia 85, 103–111. 111, 10–18. RECHER H F & DAVIS W E 2010. The foraging behaviour of WILLIAMS M R, ABBOTT I, LIDDELOW G L, VELLIOS C, WHEELER I B & woodland birds along the mulga-eucalypt line on Mt. Gibson MELLICAN A E 2001. Recovery of bird populations after Station, Western Australia during late winter and spring. clearfelling of tall eucalypt forest in Western Australia. Amytornis 2, 29–41. Journal of Applied Ecology 38, 910–920. RECHER H F, MAJER J D & GANESH S 1996. Eucalypts, WOINARSKI J C Z, RECHER H F & MAJER J D 1997. Vertebrates of and birds: on the relationship between foliar nutrients and eucalypt formations. In: Williams J E & Woinarski J C Z (eds) species richness. Forest Ecology and Management 85, 177–195. Eucalypt ecology— individuals to ecosystems, pp. 303–341. RICKLEFS R E & SCHLUTER D (Editors) 1993. Species diversity in Cambridge University Press, Cambridge ecological communities: historical and geographical WOINARSKI J C Z & TIDEMANN S C 1991. The bird fauna of a perspectives. University of Chicago Press, Chicago. deciduous woodland in the wet-dry tropics of northern SAUNDERS D A & INGHAM J 1995. Birds of southwestern Australia. Australia. Wildlife Research 18, 479–500. An atlas of changes in the distribution and abundances of the WOOLLER R D & CALVER M C 1981. Feeding segregation within wheatbelt avifauna. Surrey Beatty & Sons, Chipping Norton. an assemblage of small birds in the karri forest understorey. SERVENTY D L & WHITTELL H M 1962. Birds of Western Australia. Australian Wildlife Research 8, 401–410. Paterson Brokensha, Perth. ZAR J H 1996. Biostatistical analysis (3rd edition). Prentice Hall, SYSTAT SOFTWARE INC. 2006. SigmaStat 3.5 for Windows user’s Upper Saddle River. manual. Systat Software Inc. Point Richmond. Received 6 December 2012; accepted 10 September 2013

49 Journal of the Royal Society of Western Australia, 96(2), December 2013

Appendix 1 Acacia plot species composition, DCA site and species scores, total number of observations, species movement type and percentage of observations for each site, vegetation type and season for all acacia sites during the 2011–2012 survey period.

Site Barnong Lochada Darwin Buntine Barnong Lochada Darwin Buntine Vegetation Acacia

Season Winter Spring

DCA Axis 1 Sample Score 114 117 103 24 324 185 160 69 DCA Axis 2 Sample Score 123 97 101 237 85 105 95 227 DCA Axis 3 Sample Score 40 60 26 103 164 119 56 118 Total Number of Observations 69 125 54 61 570 147 107 74

Common name Scientific name Movement type

Australian hobby Falco longipennis Loc.Disp. ------Australian magpie Cracticus tibicen Loc.Disp. ------Australian owlet-nightjar Aegotheles cristatus Sedentary ------Australian raven Corvus coronoides Sedentary 1.45% 3.10% - 1.64% 0.18% 2.70% - 1.64% Australian ringneck Barnardius zonarius Sedentary - 6.20% 9.26% 3.28% - 0.68% 1.87% - Black honeyeater Certhionyx niger Nomadic - - - - 2.98% - - - Black-eared cuckoo Chalcites osculans Migratory - - 1.85% - - - - 3.28% Black-faced cuckoo-shrike Coracina novaehollandiae Migratory - - - 14.75% - - - - Black-faced woodswallow Artamus cinereus Nomadic ------Black-shouldered kite Elanus axillaris Loc.Disp. ------Bourke’s parrot Neopsephotus bourkii Nomadic ------Brown falcon Falco berigora Sedentary ------Brown goshawk Accipter fasciatus Loc.Disp. ------0.93% - Brown songlark cruralis Nomadic - - - - - 3.38% - - Budgerigar Melopsittacus undulatus Nomadic - - - - 76.49% 8.78% - - Chestnut-rumped thornbill Acanthiza uropygialis Loc.Disp. 4.35% 9.30% 24.07% 4.92% 0.18% 6.08% 19.63% 4.92% Cockatiel Nymphicus hollandicus Migratory - - - - 0.18% - 3.74% - Collared sparrowhawk Accipiter cirrhocephalus Sedentary ------Common bronzewing Phaps chalcoptera Loc.Disp. - - 1.85% - ---- Crested bellbird Oreoica gutturalis Sedentary 4.35% 1.55% - - 0.70% 0.68% 6.54% 6.56% Crested pigeon Ocyphaps lophotes Sedentary - - - 3.28% - - - 3.28% Crimson chat Ephthianura tricolor Nomadic - - - - - 11.49% - - Diamond dove Geopelia cuneata Nomadic ------Emu Dromaius novaehollandiae Loc.Disp. ------Galah Eolophus roseicapillus Sedentary - - 1.85% 42.62% 0.53% 8.11% - 44.26% Grey butcherbird Cracticus torquatus Sedentary ------Grey currawong Strepera versicolor Sedentary - 2.33% ------Grey fantail Rhipidura albiscapa Migratory - - 1.85% 1.64% - - 0.93% - Grey shrike-thrush Colluricincla harmonica Sedentary 2.90% 2.33% - 3.28% 0.53% 0.68% 1.87% 3.28% Horsfield’s bronze-cuckoo Chalcites basalis Migratory - - 1.85% - 0.53% - 0.93% - Inland thornbill Acanthiza apicalis Loc.Disp. 24.64% 9.30% 11.11% 3.28% 0.88% - - - Little button-quail Turnix velox Sedentary - - - - 0.88% 1.35% - 1.64% Little crow Corvus bennetti Sedentary - - - - - 6.08% - - Major Mitchell’s cockatoo Cacatua leadbeateri Sedentary ---- - 0.68% - - Malleeflowl Leipoa ocellata Sedentary - - - 1.64% - - - - Mistletoe bird Dicaeum hirundinaceum Nomadic ------0.93% - Mulga parrot Psephotus varius Sedentary - 3.10% 3.70% - - - 2.80% - Nankeen kestrel Falco cenchroides Sedentary ------Pallid cuckoo Cacomantis pallidus Migratory - - - - - 0.68% - - Pied butcherbird Cracticus nigrogularis Sedentary - 1.55% - - - 0.68% - - Pied honeyeater Certhionyx variegatus Nomadic - - - - 7.72% 8.11% 3.74% - Rainbow bee-eater Merops ornatus Migratory ------Red-capped robin Petroica goodenovii Loc.Disp. 20.29% 24.03% 25.93% 1.64% 2.46% 18.24% 14.95% 3.28% Red-tailed black-cockatoo Calyptorhynchus banksii Nomadic - 1.55% - - - 0.68% 1.87% - Redthroat Pyrrholaemus brunneus Sedentary 5.80% 6.98% 9.26% 1.64% 0.88% 1.35% 0.93% 1.64% Rufous songlark Cinclorhamphus mathewsi Migratory - - - - - 0.68% - 1.64% Rufous whistler Pachycephala rufiventris Loc.Disp. - 0.78% - - 0.53% 0.68% 8.41% 1.64% Singing honeyeater Lichenostomus virescens Nomadic 4.35% 1.55% - 9.84% 1.40% 1.35% 0.93% 16.39% Southern boobook Ninox novaeseelandiae Sedentary - - - 1.64% - - - - Southern whiteface Aphelocelphala leucopsis Sedentary 1.45% 3.10% - - - 6.76% - 3.28% Spiny-cheeked honeyeater Acanthagenys rufogularis Nomadic 1.45% - - - 0.53% 4.05% 11.21% - Splendid fairy wren Malurus splendens Loc.Disp. 14.49% 8.53% 5.56% - - 3.38% 3.74% 4.92% Striated pardalote Pardalotus striatus Migratory - 6.20% ------Variagated fairy-wren Malurus lamberti Sedentary - - - - 1.40% - - - Wedge-tailed eagle Aquila audax Sedentary ------Weebill Smicrornis brevirostris Sedentary - - 1.85% - ---- Welcome Swallow Hirundo neoxena Nomadic - - - - - 0.68% - - Western Corella Cacatua pastinator Nomadic ------1.64% Western Yellow Robin Eopsaltria griseogularis Loc.Disp. ------Whistling Kite Haliastur sphenurus Nomadic ------0.93% - White-browed Babbler Pomatostomus superciliosus Sedentary 7.25% - - 4.92% 0.53% - - 18.03% White-eared Honeyeater Lichenostomus leucotis Loc.Disp. ------1.87% - White-fronted Honeyeater Purnella albifrons Nomadic ------1.87% - White-winged Triller Lalage sueurii Migratory - 5.43% - - 0.53% 0.68% 9.35% - Willie Wagtail Rhipidura leucophrys Loc.Disp. 1.45% ------Yellow-rumped Thornbill Acanthiza chrysorrhoa Sedentary ---- - 1.35% - - Yellow-throated Miner Mamorina flavigula Loc.Disp. 5.80% ------Zebra Finch Taeniopygia guttata Nomadic ------

50 Bell et al.: Birds of the Mulga–Eucalypt Line

Barnong Lochada Darwin Buntine Barnong Lochada Darwin Buntine Acacia

Summer Autumn

295 140 109 61 119 131 87 84 100 97 118 247 147 118 129 168 114 10 16 0 66 94 28 60 DCA Species Scores 163 166 115 100 46 94 96 67 Axis 1 Axis 2 Axis 3

- - - 1.00% ------4.82% ------146 60 -346 - - 0.87% ------0.60% 2.61% 1.00% - 1.06% 1.04% 5.97% 53 58 50 - 11.45% 4.35% 2.00% - 4.26% 4.17% 5.97% 95 9 127 ------367 99 134 ------11 -39 93 - - 0.87% 1.00% ---- -132 329 -82 - - - 1.00% ---- -131 277 -167 - - - 1.00% ------4.35% - - - 29 251 172 - 3.01% ------21 -26 -176 - 1.20% ------37 -52 176 ------65.84% ------352 70 187 1.24% 22.29% 23.48% 8.00% 28.26% 18.09% 35.42% 10.45% 51 100 14 9.32% - 1.74% - ---- 293 -14 211 - 0.60% ------92 -18 -253 - 1.81% - - - 2.13% - 2.99% 35 82 -63 0.62% - 10.43% - ---- 176 122 45 - 1.81% 0.87% - - - - 5.97% -7 298 46 ------286 79 216 0.62% ------1.74% - ---- 182 54 86 - - - 4.00% - 12.77% 2.08% 13.43% 55 243 241 ---- 2.17% - 2.08% - -54 105 -47 - 3.01% ------131 40 -117 ------3.13% 1.49% -94 39 -155 1.86% 0.60% 0.87% 5.00% - - 1.04% 2.99% 151 166 131 ------264 47 -58 3.11% 2.41% 6.09% 4.00% - 3.19% - 4.48% 70 68 -8 4.97% - - 2.00% ---- 304 234 17 ------27 -108 -36 ------21.00% - - 8.33% 5.97% 38 293 -88 - 1.20% 0.87% - ---- 45 31 -183 ------253 -56 230 ---- - 4.26% - - 81 -55 222 ------328 114 153 - - 8.70% 2.00% ---- -9 230 -270 1.24% 5.42% 11.30% 3.00% 4.35% 6.38% 7.29% 7.46% 138 85 41 ------154 -59 231 9.32% 2.41% 4.35% 3.00% 4.35% - 2.08% 7.46% 220 171 -20 ------182 320 214 1.86% 2.41% 5.22% - - - 2.08% 2.99% 208 98 -4 1.24% - - 30.00% - - 1.04% 1.49% 23 367 16 ------7.85% - 19.57% 13.83% 8.33% - 131 111 138 6.83% 0.60% - 1.00% - 10.64% 3.13% - 238 54 51 11.18% 12.65% 7.85% 4.00% 21.74% 15.96% 14.58% 8.96% 188 199 90 ------74 16 198 ------0.62% ------20 -66 131 ------76 202 264 ------1.49% ------2.00% ---- 47 189 -44 ---- 6.52% - - 7.46% -4 264 17 - - - 1.00% ---- 4 230 -4 0.62% - - 1.00% ---- 263 230 -32 ------2577 173 ---- 2.17% - - 2.99% 59 235 182 - 1.81% - 2.00% - - 4.17% - -67 52 -118 ---- - 1.06% - - 122 -78 147 32.92% 18.67% - - 6.52% 6.38% - - 306 116 48

51 Journal of the Royal Society of Western Australia, 96(2), December 2013

Appendix 2 Eucalyptus plot species composition, DCA site and species scores, total number of observations, species movement type and percentage of observations for each site, vegetation type and season for all eucalypt sites during the 2011–2012 survey period.

Site Barnong Lochada Darwin Buntine Barnong Lochada Darwin Buntine Vegetation Eucalyptus

Season Winter Spring

DCA Axis 1 Sample Score 72 49 37 44 319 148 83 64 DCA Axis 2 Sample Score 83 19 22 82 66 32 99 42 DCA Axis 3 Sample Score 35 63 160 29 179 133 203 85 Total Number of Observations 150 131 88 81 922 218 247 112

Common name Scientific name Movement type

Australian magpie Cracticus tibicen Loc.Disp. ------Australian raven Corvus coronoides Sedentary 3.95% - 3.41% 2.47% 0.33% 2.78% 0.81% 9.09% Australian ringneck Barnardius zonarius Sedentary 1.32% 9.92% 1.14% - - 11.11% 4.05% 4.55% Black honeyeater Certhionyx niger Nomadic - - - - 1.19% - - - Black-eared cuckoo Chalcites osculans Migratory 1.32% - - - - - 1.21% 13.64% Black-faced cuckoo-shrike Coracina novaehollandiae Migratory - - - - 0.33% - - - Black-faced woodswallow Artamus cinereus Nomadic - - - - 0.33% - - - Bourke’s parrot Neopsephotus bourkii Nomadic ------Brown falcon Falco berigora Sedentary - 0.76% - 1.23% - - - - Brown goshawk Accipter fasciatus Loc.Disp. - - - - - 0.46% 0.81% - Brown songlark Cincloramphus cruralis Nomadic ------Budgerigar Melopsittacus undulatus Nomadic - - - - 80.80% 12.96% 7.69% - Chestnut-rumped thornbill Acanthiza uropygialis Loc.Disp. 20.39% 14.50% - 22.22% 0.98% 9.26% 0.81% 12.12% Cockatiel Nymphicus hollandicus Migratory - - - - 3.58% 6.48% 2.02% - Collared sparrowhawk Accipiter cirrhocephalus Sedentary - 0.76% ------Common bronzewing Phaps chalcoptera Loc.Disp. ------Crested bellbird Oreoica gutturalis Sedentary 1.32% 4.58% - - 0.11% 2.31% - - Crested pigeon Ocyphaps lophotes Sedentary - - - - 0.33% - - - Crimson chat Epthianura tricolor Nomadic ------Emu Dromaius novaehollandiae Loc.Disp. - - - - - 0.93% 0.40% - Galah Eolophus roseicapillus Sedentary 1.32% 1.53% 18.18% 1.23% 1.74% 2.78% 32.39% 13.64% Grey butcherbird Cracticus torquatus Sedentary 1.97% - - - 0.11% - - - Grey currawong Strepera versicolor Sedentary ------0.40% - Grey fantail Rhipidura albiscapa Migratory 4.61% 4.58% 1.14% 2.47% - - - - Grey shrike-thrush Colluricincla harmonica Sedentary 1.97% 2.29% 2.27% 1.23% 0.22% 0.93% 2.43% 6.06% Horsfield’s bronze-cuckoo Chalcites basalis Migratory - - 1.14% 1.23% - 0.46% 0.40% - Inland thornbill Acanthiza apicalis Loc.Disp. 13.82% 3.82% 5.68% 19.75% 0.11% 3.24% 5.26% 12.12% Jacky winter Microeca fascinans Loc.Disp. 0.66% - 1.14% 1.23% - - - 1.52% Little button-quail Turnix velox Sedentary ------1.52% Little eagle Hieraaetus morphnoides Sedentary ------Major Mitchell’s cockatoo Lophochroa leadbeateri Sedentary - 4.58% - - - - 0.40% - Mulga parrot Psephotus varius Sedentary 1.32% - - - 0.98% - 0.81% 1.52% Nankeen kestrel Falco cenchroides Sedentary ------Pallid cuckoo Cacomantis pallidus Migratory - - 1.14% - 0.11% 2.78% - - Pied butcherbird Cracticus nigrogularis Sedentary - - 3.41% - - 2.31% 0.40% - Rainbow bee-eater Merops ornatus Migratory ------Red wattlebird Anthochaera carunculata Nomadic - - 1.14% - - - 1.21% - Red-backed kingfisher Todiramphus pyrrhopygius Nomadic - - - - 0.33% 0.93% - - Red-capped robin Petroica goodenovii Loc.Disp. 17.76% 12.21% 5.68% 9.88% 2.17% 7.41% 2.43% 4.55% Red-tailed black-cockatoo Calyptorhynchus banksii Nomadic - - 9.09% - - 5.09% - - Redthroat Pyrrholaemus brunneus Sedentary ------4.55% Regent parrot Polytelis anthopeplus Nomadic - - 4.55% - - - 5.26% - Rufous tree-creeper Climacteris rufa Sedentary - - 4.55% - - - 2.02% - Rufous whistler Pachycephala rufiventris Loc.Disp. 0.66% 2.29% - - 0.22% - 0.81% 1.52% Singing honeyeater Lichenostomus virescens Nomadic - - - 2.47% - - - - Southern whiteface Aphelocelphala leucopsis Sedentary 12.50% 2.29% - - - 6.02% - - Spiny-cheeked honeyeater Acanthagenys rufogularis Nomadic - - - 2.47% 0.43% 2.31% - 16.67% Splendid fairy wren Malurus splendens Loc.Disp. 1.32% ------Striated pardalote Pardalotus striatus Migratory - 1.53% 2.27% - - 0.46% - - Tawny frogmouth Podargus strigoides Sedentary - - 2.27% - - - 0.40% 4.55% Tree martin Petrochelidon nigricans Migratory - - 2.27% - - - 11.74% - Varied sittella Daphoenositta chrysoptera Sedentary 2.63% ------Weebill Smicrornis brevirostris Sedentary 2.63% 29.01% 29.55% 16.05% 1.74% 12.50% 8.10% 45.45% Western corella Cacatua pastinator Nomadic - - - 2.47% - - 2.83% - Western gerygone Gerygone fusca Sedentary ------Whistling kite Haliastur sphenurus Nomadic ------1.52% White-browed babbler Pomatostomus superciliosus Sedentary 1.97% - - 8.64% - - - 3.03% White-browed treecreeper Climacteris affinis Sedentary - 1.53% ------White-eared honeyeater Lichenostomus leucotis Loc.Disp. - - - 1.23% - - - - White-fronted honeyeater Purnella albifrons Nomadic ------White-winged triller Lalage sueurii Migratory - 0.76% - - 1.84% 4.17% 2.43% 6.06% Willie wagtail Rhipidura leucophrys Loc.Disp. ------Yellow-plumed honeyeater Lichenostomus ornatus Loc.Disp. ------Yellow-rumped thornbill Acanthiza chrysorrhoa Sedentary 1.97% - - 3.70% - 0.46% - 3.03% Yellow-throated miner Mamorina flavigula Loc.Disp. 3.29% 3.05% - - 2.06% 1.85% 2.43% 3.03% Zebra finch Taeniopygia guttata Nomadic ------

52 Bell et al.: Birds of the Mulga–Eucalypt Line

Barnong Lochada Darwin Buntine Barnong Lochada Darwin Buntine Eucalyptus

Summer Autumn

165 109 0 28 41 55 36 37 101 24 0 100 130 8 86 72 55 117 88 44 21 90 193 40 DCA Species Scores 205 169 98 146 126 163 290 84 Axis 1 Axis 2 Axis 3

1.46% 2.05% - 2.74% ---- 146 60 -346 0.98% 1.37% 2.04% 2.05% 1.59% 1.84% 0.34% 1.19% 53 58 50 1.46% 15.07% 4.08% 3.42% 1.59% 9.20% 2.76% 1.19% 95 9 127 ------367 99 134 - 0.68% ------11 -39 93 1.46% - 4.08% 4.11% ---- -132 329 -82 - - - 1.37% 6.35% - - - -131 277 -167 ---- 1.59% - - - 29 251 172 - - 2.04% - - 0.61% - - 21 -26 -176 - 0.68% 4.08% - ---- 37 -52 176 - 0.68% ------2.44% ------352 70 187 10.24% 8.22% 9.18% 7.53% 33.33% 17.18% 10.00% 19.05% 51 100 14 5.85% 16.44% - - - - 2.41% - 293 -14 211 ------92 -18 -253 - 3.42% - 2.05% 1.59% 0.61% - - 35 82 -63 1.95% 2.74% - 0.68% - - - 1.19% 176 122 45 0.49% 0.68% ------7 298 46 - 0.93% ------286 79 216 - 0.00% ------182 54 86 - 4.79% - 6.16% 3.97% 0.61% 24.83% 5.95% 55 243 241 ---- 1.59% 1.23% - 1.19% -54 105 -47 ---- - 0.61% 0.34% - 131 40 -117 ------0.34% 11.90% -94 39 -155 0.98% 1.37% 2.04% 2.05% - - 1.72% 1.19% 151 166 131 ------264 47 -58 6.83% 2.74% 1.02% 7.53% 2.38% 1.84% 0.69% 7.14% 70 68 -8 ------82 -64 -61 0.98% - - 1.37% ---- 304 234 17 - - - 2.05% ------1.23% - - 27 -108 -36 3.41% - - - 10.32% - - 4.76% 38 293 -88 - - 1.02% - ---- 45 31 -183 ------253 -56 230 - 8.22% 4.08% - 0.79% 1.84% 2.41% - 81 -55 222 - 1.37% - 3.42% ---- 328 114 153 - - 1.02% 0.68% ---- -9 230 -270 - 0.68% ------230 -114 219 4.88% 0.00% 1.02% 1.37% 11.90% 4.91% 3.79% 7.14% 138 85 41 - 1.37% - - - - 1.38% - 154 -59 231 2.44% - 3.06% - - 1.23% - 2.38% 220 171 -20 ------83 -58 295 - - 10.20% - - - 3.45% - 182 320 214 1.95% - 1.02% 1.37% - 0.61% - 1.19% 208 98 -4 1.46% - - 6.85% - 1.84% 2.41% - 23 367 16 5.85% - - - 3.17% 5.52% - - 131 111 138 15.12% - - 3.42% - - 0.34% - 238 54 51 4.88% 6.85% - - 2.38% - - 3.57% 188 199 90 - - 1.02% 3.42% - - 3.10% - -74 16 198 ------61 -127 101 ------21.03% - 19 32 346 ------4.88% 19.86% 36.73% 24.66% 7.14% 30.67% 15.17% 22.62% -20 -66 131 ------76 202 264 0.49% ------0.68% ------47 189 -44 1.46% - - 6.16% 3.97% - - 3.57% -4 264 17 ---- - 1.23% - - -117 -148 -46 - - - 2.05% - - 1.03% - 4 230 -4 2.93% ------263 230 -32 ------2577 173 1.95% - - - 2.38% - 0.69% - 59 235 182 - 3.42% ------0.49% - 9.18% 2.05% 3.97% - 1.38% - -67 52 -118 1.46% 9.59% 3.06% 1.37% - 17.18% 0.34% 4.76% 122 -78 147 10.73% 2.74% ------306 116 48

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